Seasonal effects on the feeding ecology and habitat of Chersina Angulata in the South Western Cape

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Nearly one-third of the world’s tortoises live in South Africa, but little is known about their habitat requirements and feeding ecology. Chersina angulata, the angulate tortoise, is endemic to southern Africa, with a wide distribution along the western and southern coasts. Because this tortoise occupies a number of different habitat types, it has always been considered a generalist herbivore, although little is known about its diet and other needs. This study evaluates the habitat characteristics and feeding ecology of C. angulata at two study sites in the southwestern Cape, the West Coast National Park (WCNP) and Dassen Island (DI). The WCNP is a large conserved area in the Fynbos biome, along the southwestern coast of South Africa, whereas DI is a small offshore island with low floral and faunal diversity, just south of the WCNP. The efficacy of three methods used to study the feeding ecology of herbivores, focal observations, macroscopic faecal analysis and histological analysis of scats, was evaluated. Plant cover, species diversity, and the variety of growth forms were substantially larger at the WCNP than on DI. In the WCNP, shrubs and grasses were the dominant growth forms but the vegetation also included herbs, succulents, restios, sedges and parasitic plants. A few perennial species such as the grass Ehrharta villosa, shrubs such as Helichrysum niveum, Nylandtia spinosa and Rhus spp., and succulents such as Carpobrotus edulis and Ruschia spp., provided most of the plant cover. DI had a depauperate flora, consisting of succulents and herbs, and ephemeral plants contributed more than perennials did to plant cover throughout the year. The succulents Mesembryanthemum crystallinum and Tetragonia fruticosa provided most of the cover on DI. Angulate tortoises are herbivores and 72 diet plants in 32 plant families were identified to the species or genus level. Several diet species, however, could not be identified. In addition to angiosperms, the tortoises’ diet included mosses, mushrooms, insects,snails and animal faeces. The most important growth forms in the diet were herbs and grasses. The diet of the WCNP tortoises was more diverse than the diet of DI tortoises, but the number of principal food items in the diet did not differ between the two sites. Over an annual cycle, WCNP tortoises had four principal food plants while DI tortoises had five principal food plants. At both sites, principal food plants changed with the season and few plants remained principal food items in more than one season. Cynodon dactylon was a principal food item in three of the four seasons in the WCNP, whereas Trachyandra divaricata was a principal food plant each season on DI. Most principal food plants were grass or herb species but the sedge Ficinia nigrescens, and a succulent that could be identified only to the family level (Aizoaceae), featured strongly in the spring diets of DI and WCNP tortoises, respectively. The three study methods did not provide the same type or quality of information about the feeding ecology of angulate tortoises. The small size and wary nature of angulate tortoises compromised focal studies because it was often not possible to see what the tortoises ate. This method, however, provided the interesting observation that rabbit faecal pellets contributed nearly 30% to summer and autumn diets on DI when food was scarce. Rabbit faeces may not only provide a source of nutrients but may also supplement the microflora, required to digest cellulose, in the tortoises’ guts. Macroscopic evaluation of the tortoises’ scats appeared to be an ineffective method to identify diet plants, and the bulk of the scat mass could not be identified. This indicates that angulate tortoises either selected food low in fibrous content or that the digestive system of the tortoises dealt efficiently with tough plant material. The macroscopic method was the only method that highlighted the large contribution of fruits / seeds to the diet of angulate tortoises. Since the tortoises digested many seeds only partially, or not at all, C. angulata is potentially an important agent of seed dispersal in the southwestern Cape. The macroscopic study showed that on DI, sand made up 28% of the scat mass in spring, whereas sand never made a substantial contribution to the scat composition of WCNP tortoises. Lithophagy may be an important strategy in a depauperate habitat, such as DI, because the abrasive action of sand may help with the digestion of tough plants, or the sand may provide the tortoises with important minerals that are deficient in their food plants.The histological analysis of scats provided the most comprehensive diet list for C. angulata. Selection indices based on data from the histological analysis indicated that angulate tortoises were highly selective in their food choice. Most of the principal food items were selected out of proportion to their availability and the tortoises avoided the most abundant plants in their habitats. Several factors, such as palatability, accessibility and profitability, may have influenced their food choice. The proportional similarity indices for WCNP and DI tortoises, respectively, were 0.31 and 0.16, confirming that C. angulata is a food specialist and not a food generalist as was previously thought. This factor should be considered in the management of this species and in future conservation planning of its habitat.